wood preference of selected malaysian subterranean termites

Wood Preference of Selected Malaysian Subterranean Termites (Isoptera: Rhinotermitidae, Termitidae)

Peng-Soon Ngee1.2, Ai Tashiro3, Tsuyoshi Yoshimura3, Zairi Jaall & Chow-Yang

ABSTRACT

Laboratory preference of 15 Malaysian wood species: nyatoh (Ganua sp.), jelutong (Dyera costulata), teak (Tectona grandis), angsana (Pterocarpus indicus) , pine (Pinus caribaea) , balau [Shorea sp.) , red meranti (Shorea sp.), white meranti (Shorea sp.), rubber (Hevea brasieliensis), merbau (Intsia palembanica), membatu (Parinari sp.), terentang (Campnosperma auriculata) , medang (a species of Lauraceae) , melunak [Pentace sp.) and perah (Elateriospermurn tapos) and 14 Japanese wood species: hiba (Thujopsis dolabmta) , natural hinoki (Charnaecyparis obtusa), planted hinoki (Charnaecyparis obtusa), ezomatsu (Picea jezoensis), todomatsu (Abies sachaliensis), katsura [Cercidiphyllurnjaponicurn), kuri [Castanea crenata], sugi (Cryptorneria japonica), keyaki (Zelkova serrata), kusunoki (Cinnarnornun carnphora) , mizunara (Quercus rnongolica) , karamatsu (Larix leptolepis) , akamatsu (Pinus densijZora), buna (Fagus crenata) and akagashi (Quercus acuta) were evaluated against selected Malaysian termite pest species using choice-feeding tests. Results indicated that rubber, jelutong and terentang were the most preferred species among the 29 wood species tested in the laboratory on four Malaysian termite species (Coptotermes gestroi [Wasmannl , Coptotermes curvignathus Holmgren, Globitermes sulphureus (Haviland) and Microcerotermes crassus Snyder). Field studies of 15 Malaysian wood species on mixed termite species, and on Macrotermes gilvus [Hagen) confirmed the findings from the laboratory evaluation.

Keywords: Wood preference, Malaysia, Coptotermes gestroi, Coptoterrnes curvignathus, Microceroterrnes crassus, Globitermes sulphureus, Macrotennes gilvus

'Urban Entomology Laboratory. Vector Control Research Unit, School of Biological Sciences, Universiti Sains Malaysia, 11800 Penang, MALAYSIA 'Present address: Kiwi Manufacturing Sdn Bhd., Homesafe Products (M) Sdn Bhd, 8, Jalan Hasil, Kawasan Tampoi, 81200 Johor Bahru, Johor, MALAYSIA 3Laboratory of Deteroriation Control. Wood Research Institute. Kyoto University, Uji, Kyoto 61 1-001 1, JAPAN 4Reprint requests should be addressed to: Email: [email protected]

536 Sociobiology Vol. 43, No. 3, 2004

INTRODUCTION

The natural environment contains many wood species that can be utilized by termites as food. However, certain wood species are more preferred than the others. Generally, termite feeding activity is influ- enced by its density (Becker 1969, Sornnuwat et al. 1995), wood hardness (Behr et al. 19721, moisture in wood (Delaplane & La Fage 1989a), damage done by conspecifics (Delaplane & La Fage 1989b1, wood extractives (Carter 1979, Carter & Beal 1982, Carter et al. 1983, Su & Tamashiro 1986, Waller 1989, Sornnuwat et (11. 19951, wood decayed by fungus (Smythe et al. 1971), soldier proportions (Su & La Fage 1987). caste composition (Watson et al. 1978), temperature (Harverty & Nutting 1974, Smith & Rust 1993a) and colony variation (Creffield et al. 1985).

Smythe & Carter (1970a) and Morales-Ramos & Rojas (2001) reported the importance of wood combination offered to termites when doing laboratory choice feeding tests. It was shown that the preference of termites to a particular wood species could be altered by the wood combination offered to them. Smythe & Carter (1970b) tested the feeding preference of R. virginicus on 1 1 North American wood species and found that under the choice feeding tests, termite preference of four wood species increased, when compared to that in force feeding tests. This indicated that the choice feeding test was the more appropriate method to be used in determining termite wood preference than the non-choice test (force-feeding) because under the latter test method, termites were forced to feed on whatever resource available for survivor- ship.

Earlier, it was reported that several pendomestic species such as Globitermes sulphureus, Microcerotermes spp. and Macroctermes gilvus did not respond well to paper-based bait matrix (Lee 2002a, 2002b). We speculated that this could be due to the nature of matrix which is not preferred by these termite species. Thus, we hope that by determining the wood preference of Malaysian subterranean termite species, it would be possible to find a suitable wood species that can be used as part of a bait matrix against a wide spectrum of termite pest species. In this study, we tested the laboratory preference of various commercially available Malaysian wood species against four species of Malaysian subterranean termites (Coptotennes gestroi 13 colonies], Coptotermes curvignathus , Microcerotermes crassus and Globitermes sulphureus. Laboratory preference of these termite species for Japanese wood species was also evaluated for comparison purposes. Field preference studies were conducted using Malaysian wood species against mixed

P.-S. Ngee eta/. - Wood Preferences of Malaysian Subterranean Termites 537

termite species in a natural environment, a s well a s against Macrotermes gilvus.

MATERlALS AND METHODS

Laboratory preference test Test insects Coptotermesgestroi(three colonies), C. curuignathus and G. sulphureus

were collected from infested rubber (Hevea brasiliensis) wood stakes in underground monitoring stations (40 x 25 x 15 cm) on Penang Island, Malaysia. As for M. crassus, direct excavation of their aboreal nests was done. The collected termites were brought back to the laboratory and separated from debris using a slightly modified method as described by Tarnashiro et al. (1973).

Wood species A total of 29 wood species (untreated) from Malaysia and Japan were

tested namely: Malaysia - nyatoh (Ganua sp.) , jelutong (Dyera costukrta) , teak (Tectona grandis), angsana (Pterocarpus indicus) , pine (Pinus caribaea), balau (Shorea sp.), red meranti (Shorea sp.), white meranti (Shorea sp.), rubber (Hevea brasieliensis), merbau (Intsiapalembanica), membatu [Parinari sp .) , terentang [Campnosperma auriculata) , medang (a species of Lauraceae), melunak (Pentace sp.) and perah (Elateriospermum tapos); Japan - hiba (Thujopsis dolabmta), natural hinoki (Chamaecyparis obtusa), planted hinoki (Chamaecyparis obtusa), ezomatsu (Picea jezoensis), todomatsu (Abies sachaliensis), katsura (Cercidiphy llum japonicum) , kuri (Castanea crenata) , sugi {Cryptomeria japonica) , keyaki (Zelkova serrata) , kusunoki (Cinnamornun carnphora) , mizunara (Quercus mongolica), karamatsu ( h r i x leptolepis), akarnatsu (Pinus denstfira), buna (Fagus crenata) and akagashi (Quercus acuta). All Japanese wood used in this study were from the heartwood.

Experimental procedures A choice test was executed to study wood preference. A rectangular-

shaped polyethylene container (40 cm x 25 cm x 15 cm) was used as test arena for this study. Three hundred g of moistened water-washed sand (mesh size 40) was placed evenly within the container. All wood species described above in pre-weighed wood blocks (2 x 2 x 1 cm) were oven- dried, and placed randomly in the container in the order of 6 x 5. Approximately 5000 workers and 250 soldiers were introduced equally (non-localized) into the container and the experiment was replicated 10 times for each of the termite species tested. The containers were left in total darkness for 30 days under environmental conditions of 26.0 2 2.0 "C and 60 + 5 % relative humidity.


After this period, all the wood blocks were recovered, washed, oven- dried at 80 O C and weighed to determine weight loss. Wood consumption in grams and the percentage of each wood species was calculated.

Field wood preference test against mixed termite species Test site The location of the field test was Lurah Burung, a small secondary

forest reserve in Universiti Sains Malaysia Minden campus. Our earlier observation found that there were various termite species from several genera including Coptotermes, Globitermes, Microtermes, Odontotermes, Macrotermes, Subuliolitermes, Pericapritermes and Microcerotermes in the test location.

Experimental procedures Cylindrical-shaped containers (25 cm diameter x 15 cm height) were

used in this study. Fifteen Malaysian wood species measuring 10 cm x 1.5 cm x 1.5 cm (4 stakes in one group) were oven-dried, pre-weighed and placed inside the container in a random design. A total of 20 replicates were prepared and inserted completely into the soil. Each replicate was set at least 2 m apart from each other. The experiment was terminated after 3 months. All wood samples were brought back to the laboratory, washed, oven-dried and weighed. One important parameter was recorded, i.e. total termite-contact, which was based on either one of the three criteria: termite feeding, deposited faecal material or mud and gallery built on the wood. This was expressed in percentage of contact (96). All wood blocks were also visually rated based on a modified rating system of the original American Standards for Testing and Materials [ASTM 1984) method: 0 = no attack; 1 = slight superficial attack; 2 = superficial to medium attack (not deep inside); 3 = heavy attack (to penetration); 4 = very heavy attack (almost collapsed) to completely consumed.

Field wood preference test against M. giluus Test site The test was conducted near the Sports Complex of Universiti Sains

Malaysia Minden campus where many active M. gilvus mounds were located.

Experimental procedures Cylindrical-shaped containers (25 cm diameter x 15 cm height) were

used in this study. Fifteen Malaysian wood species were prepared in blocks of 10 x 1.5 x 1.5 cm (2 stakes in one group), oven-dried, pre- weighed and placed inside the container in randomized manner. A total of nine replicates were prepared and three each were inserted into a M. gilvus mound for a period of two months. Upon termination of the

P.-S. Ngee et al. - Wood Preferences of Malaysian Subterranean Termites 539

experiment, mean wood consumption of termites for each wood species was presented in gram mass loss.

Data analyses Data in mass loss (g) were subjected to analysis ofvariance, and their

means were separated using Tukey's HSD. Data in percentage were transformed to arc-sine values before analysis of variance.

RESULTS AND DISCUSSION

Laboratory preference test Choice feeding test indicated that rubber wood was the most

Table 1: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against C. gestroi (colony A).

Wood species Common Mean wood mass loss Mean wood mass loss name + SEMI (g) + SEMI (%)

Shorea sp. Balau 0.0141 + 0.0024 a 0.4338 + 0.0849 a Pterocarpus indicus Angsana 0.0199 + 0.0020 a 0.7388 + 0.0587 ab Chamaecyparis obtuse Hinoki (planted) 0.0220 + 0.0025 a 1.4091 + 0.1 144 abc Chamaecyparis obtuse Hinoki (natural) 0.0261 + 0.001 9 a 1.2297 + 0.1405 abc Ctyptomeria japonica Sugi 0.0304 + 0.0046 a 2.1071 + 0.3136 a-d Quercus acuta Akagashi 0.0329 + 0.0037 a 1.1834 + 0.1373 abc Abies sachaliensis Todomatsu 0.0436 + 0.0053 a 2.9254 + 0.3980 a-d Species of Lauraceae Medang 0.0441 + 0.0055 a 1.3705 + 0.1844 abc Castanea crenata Kuri 0.051 1 + 0.0076 a 2.8292 + 0.4395 a-d Tectona grandis Teak 0.0605 + 0.01 18 ab 2.6663 + 0.1745 a-d Pinus densiflora Akamatsu 0.0740 + 0.0099 ab 3.5999 + 0.4562 a-e Zelkova serrata Keyaki 0.0869 + 0.01 I I abc 2.9442 + 0.3461 a-d lntsia palembanica Merbau 0.1317 + 0.0168 a-d 3.3788 +_ 0.4390 a-d Shorea sp. White meranti 0.1487 k 0.0201 a-e 3.8255 + 0.5959 a-e Quercus mongolica Mizunara 0.1612 + 0.0282 a-e 6.1088 + 0.7944 a-e Parinari sp. Membatu 0.1650 + 0.0203 a-e 4.4210 + 0.5567 a-e Cercidiphyllum japonicum Katsura 0.1677 + 0.0019 a-f 9.5195 + 1.0499 def Picea jezoensis Ezomatsu 0.1684 + 0.0221 a-f 10.7377 + 1.4133 ef Thujopsis dolabrata Hiba 0.1690 + 0.0177 a-f 9.9456 + 0.9457 def Pentace sp. Melunak 0.2430 + 0.0348 b-f 7.9148 + 1.1943 b-e Elanteriospermum tapos Perah 0.2560 + 0.0275 c-f 8.1987 + 0.9871 cde Dyera costulata Jelutong 0.2945 + 0.0295 def 21.2314 + 2.1659 gh Larix leptolepis Karamatsu 0.3026 + 0.0263 efg 18.1306 + 1.5339 g Cinnamomun camphora Kusunoki 0.3364 + 0.0361 fg 16.3739 + 2.0815 fg Fagus crenata Buna 0.4697 + 0.0443 gh 20.4458 + 1.9755 gh Shorea sp. Red meranti 0.5098 + 0.0466 h 25.7219 + 2.5729 h Campnosperma auriculata Terentang 0.5875 + 0.0460 hi 46.0207 + 4.0566 i Ganua sp. Nyatoh 0.61 97 + 0.0786 hi 15.7470 + 1.9212 fg Pinus caribaea Pine 0.7543 + 0.0486 i 44.0792 + 2.3390 i Hevea brasieliensis Rubber 1.2099 + 0.0917 j 45.0819 + 2.3742 i

'Means within the same column followed by different letters are significantly different (Tukey HSD, p < 0.05).


preferred wood among the 30 wood species when tested against C. gestroi colony A (Table 1). Mean wood mass losses showed that five Malaysian woods namely rubber, pine, nyatoh, terentang and red meranti (decreasing in preference) were the most preferred wood species by this colony. Differences among the latter four woods were not significant, except between pine and red meranti. Our force-feeding experiment revealed that jelutong was found to be more highly attractive than other wood species such as rubber, nyatoh, terentang and red meranti by this colony (P.S. Ngee et al., unpublished data], but when the termites were given choices, the findings became different. This reconfirmed earlier reports of interactions among the wood species Table 2: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against C. gestroi (colony B).

Wood species Common name Mean wood mass loss Mean wood mass loss k SE1 (g) + SE1 (%)

Castanea crenata Kuri Quercus acuta Akagashi Shorea sp. White meranti Pentace sp. Melunak Shorea sp. Balau Abies sachaliensis Todomatsu Pterocarpus indicus Angsana Chamaecyparis obtusa Hinoki (planted) Thujopsis dolabrata Hiba Thujopsis dolabrata Hinoki (natural) Zelkova serrata Keyaki Cryptomeria japonica Sugi Tectona grandis Teak Parinari sp. Membatu Species of Lauraceae Medang Picea jezoensis Ezomatsu Pinus densiflora Akamatsu lntsia palembanica Merbau Cercidiphyllum japonicum Katsura Shorea sp. Red meranti Elanteriospermum tapos Perah Larix leptolepis Karamatsu Dyera costulata Jelutong Quercus mongolica Mizunara Campnosperma auriculata Terentang Pinus caribaea Pine Fagus crenata Buna Ganua sp. Nyatoh Hevea brasieliensis Rubber

- - -

0.0069 k 0.0064 a 0.0165 k 0.0058 a 0.021 9 k 0.0029 a 0.0289 k 0.0070 ab 0.0301 k 0.0063 ab 0.0310 k 0.0229 ab 0.0325 k 0.0017 ab 0.0384 k 0.0017 ab 0.0396 +- 0.0059 ab 0.041 1 k 0.0034 ab 0.0537 k 0.01 39 ab 0.0597 k 0.0218 ab 0.0659 k 0.0176 ab 0.0675 k 0.0091 ab 0.0698 ? 0.0142 ab 0.0761 k 0.01 92 ab 0.0973 k 0.0296 abc 0.1019 k 0.0275 abc 0.2063 k 0.1058 a-d 0.2542 k 0.0621 a-e 0.4614 + 0.0569 a-e' 0.5540 k 0.0546 b-f 0.6213 k 0.0751 C-g 0.6418 k 0.0894 d-g 0.7642 k 0.0889 efg 0.9583 k 0.1053 fgh 1.0486 k 0.2714 fgh 1.0967 k 0.2644 gh 1.4210 k 0.1689 h

- -

0.3049 k 0.2835 a 0.5454 k 0.1889 a 0.8565 k 0.1303 a 0.9242 k 0.2285 a 0.8867 k 0.1 758 a 2.4802 k 1.8277 a 1.4342 k 0.081 1 a 2.1404 k 0.0966 a 2.1189 k 0.3141 a 2.3945 k 0.2145 a 1.7317 k 0.4501 a 4.9393 k 1.9464 a 2.7961 k 0.7486 a 1.7537 + 0.2644 a 2.8709 ? 0.7079 a 4.7684 k 1.1596 a 6.2309 k 1.8803 a 2.7026 k 0.8520 a 1 1.4374 k 5.8080 ab 13.8053 k 3.6068 abc 10.6339 k 1.9621 ab 31.9014 k 3.1186 cd 46.0002 k 5.6914 def 28.3549 k 3.8964 bcd 74.0464 k 7.0408 g 55.8489 k 6.2998 efg 38.9162 k 9.891 1 de 28.4789 k 6.4276 bcd 64.0881 k 7.4014 fg

- - - - -

'Means within the same column followed by different letters are significantly different (Tukey HSD, p < 0.05)

P.-S. Ngee et a/. - Wood Preferences of Malaysian Subterranean Termites 54 1

tested as reported by Morales-Ramos & Rojas (2001). Buna, kusunoki, kararnatsu, jelutong, perah and melunak were found to have a mean wood mass loss greater than 0.2 g, but no significant differences were detected among these wood species. The rest of the wood species showed little consumption (less than 0.2 g) by the colony and no significant differences among them were observed.

Table 2 showed the relative mean wood mass losses of 30 Malaysian and Japanese timbers during a month exposure to C. gestroi colony B. Rubber, nyatoh, buna, pine, terentang, mizunara, jelutong, karamatsu, perah, red meranti and katsura were found to be the most preferred

Table 3: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against C. gestroi (colony C).

Wood species Common name Mean wood mass Mean wood mass loss + SE1 (%) loss +SE1 (%)

-

Chamaecyparis obtusa Hinoki (planted) 0.0206 & 0.0016 a 1.1613 + 0.0880 a Ctyptomeria japonica Sugi 0.0243 & 0.0012 a 1.8518 & 0.0767 ab Thujopsis dolabrata Hiba 0.0269 + 0.0047 a 1.5418 + 0.2829 ab Abies sachaliensis Todomatsu 0.0303 + 0.01 16 a 2.0769 + 0.7282 ab Quercus acuta Akagashi 0.031 1 + 0.0074 a 1.0850 + 0.2650 a Shorea sp. Balau 0.0322 + 0.0034 a 0.9348 + 0.0920 a Castanea crenata Kuri 0.0323 + 0.0015 a 1.6159 + 0.1088 ab Chamaecyparis obtusa Hinoki (natural) 0.0346 + 0.001 1 a 1.9881 + 0.0553 ab Species of Lauraceae Medang 0.0346 + 0.0070 a 0.9738 + 0.1939 a Pterocarpus indicus Angsana 0.0403 + 0.0035 a 1.6910 + 0.1057 ab Pinus densiflora Akamatsu 0.0404 + 0.0043 a 2.3422 ? 0.2706 ab Picea jezoensis Ezomatsu 0.0408 & 0.0064 a 2.5791 + 0.3931 ab Pentace sp. Melunak 0.0458 + 0.0096 a 1.4573 + 0.3041 ab Zelkova serrata Keyaki 0.0614 + 0.0161 a 2.0339 + 0.5224 ab Parinari sp. Membatu 0.0662 + 0.0089 a 1.7048 + 0.2293 ab Tectona grandis Teak 0.0669 + 0.0182 a 2.831 1 + 0.7675 ab Shorea sp. White meranti 0.0745 + 0.0227 a 2.2833 + 0.6811 ab lntsia palembanica Merbau 0.0818 + 0.0125 a 2.2298 + 0.3579 ab Cercidiphyllum japonicum Katsura 0.0843 + 0.0226 a 4.7951 + 1.2577 abc Cinnamomun camphora Kusunoki 0.1078 + 0.0156 a 4.6379 + 0.5425 abc Larix leptolepis Karamatsu 0.2180 i 0.0300 ab 12.6508 i 1.7523 a-d Shorea sp. Red meranti 0.2480 & 0.0905 ab 11.6219 + 4.21 14 a-d Ganua sp. Nyatoh 0.2567 + 0.0391 ab 6.8324 + 0.9690 abc Dyera costulata Jelutong 0.2679 2 0.0134 ab 20.1707 + 1.1433 cd Hevea brasieliensis Rubber 0.5377 + 0.0673 bc 16.9615 + 2.0239 bcd Fagus crenata Buna 0.5819 + 0.0853 bc 23.8701 + 3.5656 de Campnosperma auriculata Terentang 0.5984 + 0.0496 bc 55.7917 + 6.9437 f Elanteriospermum tapos Perah 0.7091 + 0.3139 cd 16.7610 + 7.4055 bcd Quercus mongolica Mizunara 0.8819 & 0.1806 cd 37.9501 + 8.3505 e Pinus caribaea Pine 1.0212 & 0.1086 d 58.6015 + 6.7721 f



wood species. Kuri, akagashi, white meranti, melunak, balau, todomatsu, angsana, hinoki (both natural growth and planted), hiba, keyaki, sugi, teak, membatu, medang, ezomatsu, akarnatsu and merbau were the least attacked wood species with an average wood mass losses of < 0.2 g and no significant differences were found among these wood species. The results obtained were comparable to those with C. gestroi colony A, where the similar group of wood species was also identified as non-preferred ones.

Similar results for non-preferred species were also recorded for colony C of C. gestroi (Table 3). However, there appeared to be some variations for the most preferred species. Pine was the most preferred

Table 4: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against C. curvignathus..

Wood species Common name Mean wood mass Mean wood mass loss + SE1 (g) loss + SE1 (%)

Zelkova serrata Shorea sp. Pentace sp. Abies sachaliensis Shorea sp. Chamaecyparis obtusa Quercus acuta Thujopsis dolabrata Pterocarpus indicus Castanea crenata Parinari sp. Tectona grandis Chamaecyparis obtusa Species of Lauraceae Pinus densiflora lntsia palembanica Cercidiphyllum japonicum Picea jezoensis Cryptomeria japonica Elanteriospermum tapos Shorea sp. Larix leptolepis Hevea brasieliensis Ganua sp. Fagus crenata Campnosperma auriculata Dyera costulata Pinus caribaea Quercus mongolica

-

Keyaki Balau Melunak Todomatsu White meranti Hinoki (natural) Akagashi Hiba Angsana Kuri Membatu Teak Hinoki (planted) Medang Akamatsu Merbau Katsura Ezomatsu Sugi Perah Red meranti Karamatsu Rubber Nyatoh Buna Terentang Jelutong Pine Mizunara

-

0.6863 + 0.1741 a 1.1816 + 0.3584 a 1.5243 + 0.301 9 a 4.7643 + 1.5522 abc 1.6909 + 0.6079 a 3.8503 + 2.3649 abc 2.7188 + 0.8126 ab 4.6148 + 1.8186 abc 3.7945 f 1.2489 abc 5.7320 + 1.7699 abc 3.1467 f 0.9522 ab 5.2617 + 2.2629 abc 7.1499 + 3.9795 abc 5.8150 + 2.3881 abc 9.3933 + 1.9250 abc 5.6346 + 1.6509 abc

15.3060 + 3.3475 a-d 19.3758 + 3.1945 bcd 27.5647 + 3.4372 def 14.6272 f 3.9613 a-d 36.8936 + 5.4654 ef 39.7592 + 4.4698 f 30.1955 + 4.1064 def 20.6879 k 3.7463 cde 29.5325 + 4.4560 def 80.4927 + 1.5332 h 72.9861 + 7.6281 gh 74.3674 + 4.3261 gh 60.3724 + 6.6409 g

'Means within the same column followed by different letters are significantly different (Tukey HSD; p < 0.05)

P.-S. Ngee et a/. - Wood Preferences of Malaysian Subterranean Termites 543

wood, not rubber as observed for colony A & B. No significant difference was noticed among mizunara, perah, terentang, buna, rubber, jelutong, nyatoh, red meranti and karamatsu, except between mizunara and jelutong, nyatoh, red meranti, karamatsu.

C. curvignathus, the most aggressive termite species, showed a great preference for mizunara and pine, but no significant difference was found between the two species [Table 4). This was followed by jelutong, terentang, buna, nyatoh, rubber, kararnatsu, red meranti and perah with no significant difference observed among them [with exception to between jelutong, red meranti and perah). Generally, mean mass losses

Table 5: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against G. sulphureus..

Wood species Common name Mean wood mass Mean wood mass loss t SE1 (g) Loss k SE1 (%)

Thujopsis dolabrata Hiba 0.0149 f 0.0005 a 0.0891 f 0.0289 a Picea jezoensis Ezomatsu 0.0183 k 0.0050 a 1.171 1 ? 0.3686 a Abies sachaliensis Todomatsu 0.0218 k 0.0023 a 1.5079 t 0.1609 a Chamaecyparis obtusa Hinoki (planted) 0.0312 k 0.0107 ab 1.6883 k 0.6091 a Tectona grandis Teak 0.0315 k 0.0055 ab 1.4457 k 0.2237 a Chamaecyparis obtusa Hinoki (natural) 0.0337 k 0.0009 ab 1.8274 k 0.021 1 ab Pterocarpus indicus Angsana 0.0338 k 0.0050 ab 1.3876 + 0.1932 a Shorea sp. Balau 0.0366 k 0.0046 ab 1.0341 ? 0.1316 a Pinus densiflora Akamatsu 0.0418 k 0.0009 ab 2.0110 k 0.0251 abc Pinus caribaea Pine 0.0432 k 0.0094 ab 3.1246 k 0.5615 abc Quercus acuta Akagashi 0.0441 k 0.0038 ab 1.6056 k 0.131 1 a Clyptomeria japonica Sugi 0.0574 k 0.0189 abc 3.9463 k 1.3050 a-d Larix leptolepis Karamatsu 0.0577 k 0.0153 abc 3.4202 f 0.7783 abc Zelkova serrata Keyaki 0.0623 f 0.0010 abc 2.3165 k 0.0329 abc Parinari sp. Membatu 0.0631 t 0.0040 abc 1.6927 k 0.1 195 a Species of Lauraceae Medang 0.0669 k 0.0108 abc 2.0415 k 0.3248 abc Cinnamomun camphora Kusunoki 0.0800 k 0.01 12 abc 4.0898 k 0.5927 a-d lntsia palembanica Merbau 0.1 102 k 0.0250 a-d 2.9376 t 0.6394 abc Ganua sp. Nyatoh 0.1 166 k 0.0056 a-d 3.0241 k 0.1883 abc Pentace sp. Melunak 0.1 172 k 0.0280 a-d 3.7322 k 0.8806 abc Shorea sp. White meranti 0.1365 k 0.0235 a-d 3.6949 k 0.661 1 abc Shorea sp. Red meranti 0.1393 ? 0.0297 a-d 8.1669 k 1.8881 b-f Dyera costulata Jelutong 0.1 524 k 0.0250 a-d 11.1392 k 2.0071 ef Campnospema auriculata Terentang 0.1670 k 0.0240 bcd 11.6288 k 2.1028 ef Quercus mongolica Mizunara 0.1887 k 0.0234 cd 7.2786 f 0.9216 a-f Cercidiphyllum japonicum Katsura 0.1899 k 0.0659 cd 10.6590 f 3.6323 ef Elanteriospemum tapos Perah 0.1906 k 0.0701 cd 5.2317 k 1.6997 a-e Hevea brasieliensis Rubber 0.2290 k 0.0822 d 8.3476 k 2.9846 c-f Fagus crenata Buna 0.2356 k 0.0214 d 10.2909 k 0.9493 def Castanea crenata Kuri 0.2386 t 0.0570 d 13.4460 k 3.2379 f



of wood due to attack by C. curuignathus were more clearly differenti- ated than those fed by C. gestroi. However, its preference for the respective wood species were generally similar to those recorded for C . gestroi.

It is interesting to note that hiba and teak, two highly resistant wood species and least preferred by Japanese Coptotermes fornosanus (Imamura 200 l ) , was found to be moderately preferred by the Malay- sian Coptotermes spp. used in this study. On the other hand, buna which was classified to be moderately resistant to C. fonnosanus (Imamura 200 1) was highly preferred by Malaysian Coptotermes species with mean wood mass loss between 20 - 39% after 30 days choice-

Table 6: Mean mass loss of various Malaysian and Japanese wood species after 30 days choice- feeding test against M. crassus..

Wood species Common name

Thujopsis dolabrata Cryptomeria japonica Shorea sp. Pentace sp. Abies sachaliensb Pinus densiflora Chamaecyparis obtuse Parinari sp. Larix leptolepis Pterocarpus indicus Chamaecyparis obtusa Tectona grandis Cercidiphyllum japonicum Quercus acuta Castanea crenata Species of Lauraceae Shorea sp. Pinus caribaea Zelkova serrata lntsia palembanica Shorea sp. Elanteriospermum tapos Dyera costulata Fagus crenata Picea jezoensis Ganua sp. Quercus mongolica Hevea brasieliensis Campnospenna auriculata

Hiba Sugi Balau Melunak Todomatsu Akamatsu Hinoki (natural) Membatu Kararnatsu Angsana Hinoki (planted) Teak Katsura Akagashi Kuri Medang Red rneranti Pine Keyaki Merbau White meranti Perah Jelutong Buna Ezomatsu Nyatoh Mizunara Rubber Terentang

Mean wood mass loss + SET (g)

0.0144 + 0.0004 a 0.0236 + 0.0017 ab 0.0255 + 0.0022 ab 0.0266 + 0.0017 ab 0.0271 + 0.0013 ab 0.0318 + 0.0017 ab 0.0342 i: 0.0036 ab 0.0369 + 0.0016 ab 0.0369 + 0.0099 ab 0.0395 + 0.0024 ab 0.0481 + 0.0082 abc 0.0487 + 0.0067 abc 0.0656 + 0.0128 a-d 0.0716 + 0.0183 a-d 0.0882 + 0.0148 a-d 0.0923 + 0.0362 a-d 0.1076 + 0.0244 a-e 0.1 132 + 0.0366 a-e 0.121 1 f 0.0166 a-e 0.1362 2 0.0199 a-e 0.1 522 + 0.0262 a-e 0.1560 + 0.0281 b-e 0.1577 2 0.0130 b-e 0.1587 & 0.031 3 b-f 0.1880 + 0.0330 C-f 0.1949 + 0.0532 def 0.2397 + 0.0288 ef 0.2987 + 0.051 9 f 0.7126 + 0.0873 g

Mean wood mass loss + SET (%)

0.8496 + 0.0243 a 1.6171 + 0.1109 a 0.7433 + 0.0573 a 0.8535 + 0.0539 a 1.8569 + 0.0879 a 1.5171 + 0.0782 a 1.8586 + 0.1926 a 0.9478 + 0.0451 a 2.1823 + 0.5881 a 1.4030 + 0.081 1 a 2.5475 + 0.3484 a 2.1 130 + 0.2986 a 3.7020 + 0.6961 a 2.5608 + 0.6604 a 4.9051 + 0.8277 a 2.6006 + 0.9372 a 5.0240 + 1.0211 a 6.771 1 * 2.1896 a 4.1461 + 0.5472 a 3.4940 f 0.5523 a 3.8389 + 0.7264 a 4.9159 + 1.1101 a 11.4436 + 0.9751 a 6.8613 + 2.8284 a

11.7724 + 2.0375 a 5.0862 & 1.4251 a 8.9087 + 1.0729 a 9.9480 + 1.7035 a

66.0917 + 11.0652 b



feeding test. This suggested a differential feeding preference among different species of Coptotermes toward wood species that were uncom- mon to the foraging colonies.

G. sulphureus, a higher termite species, is a slow feeder [Ngee & Lee 2002, Lee et al. 2003). It was found to share the same feeding preference as Coptotermes spp., but its consumption rate on preferred wood species was much lower (Table 5). Of the 30 wood species offered to this species, kuri, buna and rubber were among the three most preferred. However, it is not possible to separate the wood species into preferred and non-preferred groups due to lack of significant difference among them.

On the other hand, M. crassus demonstrated a clear difference in their feeding patterns against the tested wood species. This species was found to only prefer only terentang, and showed a relatively poor response on other wood species (Table 6). Approximately 66.09% of the terentang wood block was consumed at the end of the experiment. It was anticipated that if the experimental period was extended, different feeding patterns may be detected upon terentang wood supply in the arena diminished. This species may have a slower exploratory behavior, thus more time was needed to explore the other potential wood species. This factor, however cannot be proven in this study, unless a longer evaluation period was done.

Field wood preference test against mixed termite species Field evaluation of 15 Malaysian wood species to subterranean

termite attacks showed that significantly higher mean wood mass loss for rubber when compared to other wood species (Table 7). This pointed the suitability of using rubber as a termite bait matrix. Pine was ranked as the second most preferred wood after rubber. Visual rating demonstrated relatively comparable results to the mean wood mass losses where both species suffered "heavy" to "very heavy" attack by various termite species in the field. On the other hand, balau, white meranti, medang, melunak, merbau, membatu and teak registered "no at tack to "slight superficial at tack, whereas red meranti, angsana, nyatoh and perah were rated to have "slight superficial" to "medium attack.

Field wood preference test against mixed termite species M. gilvus has been reported to be a peridomestic termite pest species

in Malaysia that may cause infestation to buildings and structures upon elimination of the more dominant Coptotermes spp. (Lee 2002a, 2002bl. Table 8 shows the mean wood mass losses, percentage of wood contacted and visual damage assessments of 15 Malaysian wood species after two-month exposure to three field colonies of M. gilvus.

Sociobiology Vol. 43. No. 3, 2004

Results indicated that rubber was the most preferred wood ' k t h a significantly higher mean wood mass loss when compared with other wood species, followed by terentang and jelutong. No significant difference, however was detected for the latter two wood species. The remaining wood species were less preferred and were found to have no significant difference in wood mass losses. These results were also comparable with the mean visual rating for damage assessments where

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3 h8 z s 2 so?, g P, 0 n, ( lo,o!- f E. E * 9 aq 20-0 %;I s. CD (lo,

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Table 8: Mean wood consumption and wood contact of 15 Malaysian wood species by M. gilvus after 2 months exposure in the field.

Wood species Common name Mean wood Means wood Termite Mean + SE mass loss1 mass loss1 C~ntact ' ,~

(9) ( yo) (%) Visual rating

Shorea sp. White meranti - 0.0387+ 0.1939 a - 0.5064 + 0.2521 100.00 0.06 t 0.06 a Shorea sp. Balau 0.0839 + 0.0739 a 0.0959 + 0.0803 100.00 0.1 1 + 0.08 a Pentace sp. Melunak 0.0946 t 0.1016 a 0.1036 + 0.1104 100.00 0.06 + 0.06 a lntsia palembanica Merbau 0.1 149 t 0.1246 a 0.1328 + 0.1444 100.00 0.22 k 0.13 a Ganua sp. Nyatoh 0.2067 + 0.2347 a 0.3176 + 0.3629 100.00 0.1 1 + 0.1 1 a Species of Lauraceae Medang 0.4418 + 0.1133 a 0.6893 + 0.1748 100.00 0.17 + 0.09 a Pterocarpus indicus Angsana 0.7875 + 0.2014 a 1.1773 t 0.3046 100.00 0.00 + 0.00 a Parinari sp. Membatu 1.3293 + 0.4027 a 1.6280 + 0.5057 100.00 0.1 1 + 0.08 a Shorea sp. Red meranti 1.5774 + 0.8466 a 2.6790 + 1.4125 100.00 0.78 + 0.23 ab Tectona grandis Teak 2.2798 + 2.3195 a 3.1 11 1 + 3.3393 100.00 0.56 + 0.24 ab Elanteriospermum tapos Perah 3.3443 t 0.9564 a 4.8416 + 1.3994 100.00 0.83 + 0.22 ab Pinus caribaea Pine 6.5923 t 2.8233 a 13.0608 + 5.6061 100.00 2.33 + 0.30 bc Dyera costulata Jelutong 25.9405 t 4.8591 b 60.4262 k 4.8591 100.00 3.56 + 0.17 c Campnosperma auriculata Terentang 32.6265 + 3.1876 b 79.3174 + 7.1274 100.00 3.44 + 0.24 c Hevea brasieliensis Rubber 47.2344 t 8.0337 c 46.2676 + 7.9872 100.00 3.44 + 0.21 c

'Data were based on 18 stakes (9 replicates x 2 stakes per wood species) and, means within the same column followed by different letters were significantly different. 2Termite feeding, depositing fecal materials or building mud or gallery on wood. 3Visual assessment: 0 = No attack; 1 = Slight superficial attack; 2 = Superficial-medium attack (not deep inside); 3 = Heavy attack (penetrated); 4 = Very heavy attack (almost collapsed)

z nr 0) Y

I?. L1) 2


selection was based on the results obtained from various evaluations on relative preferences of these wood species to the various termite species tested. These wood species are also easily procured from most sawmills in Malaysia. Several Japanese wood species [such as hiba and buna) which were tested in this study were relatively more preferred by the Malaysian Coptotermes when compared to the earlier studies of Japanese C. fornosanus. This possibly suggested differential feeding response among Coptotermes species, a s well as towards wood species from different geographical localities.

In most commercially available baiting systems, southern yellow pine (Pinus radiate) is widely used. The choice of this species was likely based on studies which had been conducted in the United States. Our experience has revealed that rubber wood is a better choice for detection of multiple termite species in monitoring stations in Malaysia (C.Y. Lee et al., unpublished). Thus the use of a more attractive wood species may increase the percentage of monitoring devices being attacked and aggregated by termites during the monitoring process. In addition, the cost of baiting can probably be reduced if the period between installation of monitoring devices to termite attacks can be reduced. This may also decrease the cost of baiting since fewer visits to the baited site will be required of the pest control operator if the period between installation of monitoring stations and termite attack is reduced.

ACKNOWLEDGMENTS

The authors thank S.C. Lim (Forest Research Institute of Malaysia) for confirming some of the Malaysian wood species. P.-S. Ngee was supported under an M.Sc. scholarship provided by SaraLee (M) Sdn. Bhd. This work constitutes part of a collaborative research project between Wood Research Institute, Kyoto University and School of Biological Sciences, Universiti Sains Malaysia, and was partially supported by the JSPS-LIP1 Core University Program in the field of wood science.

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wood preference of selected malaysian subterranean termites

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